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Zeitschrift/Journal: Illiesia

Jahr/Year: 2008

Band/Volume: 04

Autor(en)/Author(s): Grubbs Scott A., DeWalt R. Edward

Artikel/Article: Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee (: ). 143-149

Grubbs, S.A. & R.E. DeWalt 2008. Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee Plecoptera: Perlidae). Illiesia, 4(14):143-149. Available online: http://www2.pms-lj.si/illiesia/Illiesia04-14.pdf

TAXONOMIC AND DISTRIBUTIONAL NOTES ON PERLESTA TEAYSIA, P. GOLCONDA, AND P. SHAWNEE (PLECOPTERA: PERLIDAE)

Scott A. Grubbs1 & R. Edward DeWalt2

1 Department of Biology and Center for Biodiversity Studies Western Kentucky University Bowling Green, Kentucky, U.S.A. 42101 E-mail: [email protected] 2 Illinois Natural History Survey 1816 S. Oak Street Champaign, IL 61820 USA E-mail: [email protected]

ABSTRACT Perlesta napacola DeWalt is shown to be a junior synonym of P. teaysia Kirchner & Kondratieff. The egg of P. golconda DeWalt & Stark is illustrated with scanning electron microscopy for the first time, and a significant range extension is presented and discussed for P. shawnee Grubbs.

Keywords: Plecoptera, Perlidae, Perlesta, P. teaysia, P. napacola, P. golconda, P. shawnee

INTRODUCTION Ontario (CNC), C. P. Gillette Museum of The known diversity of the eastern Nearctic Diversity, Colorado State University, Fort Collins, stonefly genus Perlesta continues to increase, as Colorado, (CSU), Illinois Natural History Survey, evidenced by the number of new species descriptions Champaign, Illinois (INHS), Ohio Biological Survey, within the last 20 years. Stark (1989) recognized 12 Columbus, Ohio (OBS), Purdue University, West species as distinct. New descriptions published prior Lafayette, Indiana (PU), and Western Kentucky to Stark’s (2004) updated taxonomic treatment, plus University, Bowling Green, Kentucky (WKU). those presented in Kondratieff et al. (2006, 2008), increased the total number of species known to 27 Perlesta teaysia Kirchner & Kondratieff (Stark et al., 2008). (Figs. 1-15) The difficulty of Perlesta resides in that multiple life stages are typically needed to identify a Perlesta teaysia Kirchner & Kondratieff, 1997:290. ♂ known species or describe a species as new to Holotype (United States National Museum of Natural History). Reed Creek of New River, Wythe Co., science. Males with fully extruded aedeagi associated Virginia, U.S.A. with females with mature eggs provide the most Perlesta teaysia Stark, 2004:95 definitive evidence of a species identity. The intent of Perlesta napacola DeWalt, 2002:174. Holotype ♂ (Illinois this paper is to use scanning electron microscopy Natural History Survey). Unnamed tributary of (SEM) of Perlesta eggs as the central means of Vermilion River, Vermilion Co., Illinois, U.S.A. New addressing taxonomic and distributional issues of P. synonymy napacola DeWalt, P. teaysia Kirchner & Kondratieff, P. golconda DeWalt & Stark, and P. shawnee Grubbs. Material examined. . U.S.A., ILLINOIS: Vermillion Specimens were examined from the Monte L. Bean Co., tributary to Vermillion River, 8 km SE Westville, Museum, Brigham Young University, Provo, Utah Forest Glen Forest Preserve, 19 June 2001, R.E. (BYU), Canadian National Collection, Ottawa, DeWalt, 1 ♂(Holotype – P. napacola; INHS); same but

Illiesia – http://www2.pms-lj.si/illiesia/ Volume 4 – Number 14 – Page 143

Grubbs, S.A. & R.E. DeWalt 2008. Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee Plecoptera: Perlidae). Illiesia, 4(14):143-149. Available online: http://www2.pms-lj.si/illiesia/Illiesia04-14.pdf

Figs. 1–15. Perlesta teaysia egg SEM micrographs. 1–3: U.S.A., IL, Vermilion Co., Forest Glen Forest Preserve, 4 August 1977, 4–6: U.S.A., IN, Jackson Co., Guthrie Creek, 18 July 2000, 7–9: U.S.A., KY, Warren Co., Doty Creek, 6 July 2003, 10–12: U.S.A., MD, Allegany Co., Flintstone Creek, 4 July 1996, 13–15: U.S.A., VA, Wythe Co., Reed Creek, 24 July 2008). 1, 4, 7, 10, 13. Entire egg, 200X; 2, 5, 8, 11, 14. Anterior pole, 1000X; 3, 6, 9, 12, 15. Chorionic sculpturing, 2000X.

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Grubbs, S.A. & R.E. DeWalt 2008. Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee Plecoptera: Perlidae). Illiesia, 4(14):143-149. Available online: http://www2.pms-lj.si/illiesia/Illiesia04-14.pdf

19 June 2001, R.E. DeWalt, 4 ♂, 7 ♀ (Paratypes – P. Kristsky, 5 ♂, 3 ♀ (OBS); Brown Co., Ellis Run Creek, napacola; INHS); same but 6 July 2000, R.E. DeWalt, 1 S.R. 41, 9 July 1994, Kovarik, 3 ♂, 8 ♀ (OBS); ♂, 1 ♀ (Paratypes – P. napacola; INHS); same but 4 Clermont Co., Little Indian Creek, 5 km SSW Laurel, July 1977, D.W. Webb, 2 ♂, 10 ♀ (Paratypes – P. 23 July 1997, G. Kristsky, 4 ♂, 2 ♀ (OBS); Clinton Co., napacola; INHS). INDIANA: Brown Co., tributary to Lytle Creek, nr. Todds Fork junction, 5 June 1944, Middle Fork Salt Creek, 3 km N Stone Head, 21 June W.E. Ricker, 2 ♂, 1 ♀ (BYU); Coshocton Co., Beaver 2008, R. E. DeWalt, 1 ♂, 2 ♀ (INHS); Bear Creek, 0.7 Run, 2 km NW Warsaw, 14 July 2001, B. Armitage, 8 km NNE Waycross, Yellowwood State Forest, 21 ♂, 3 ♀ (OBS); Simmons Run, 0.9 km SSW Warsaw, 26 June 2006, R.E. DeWalt, 1 ♂ (INHS); Floyd Co., June 2000, E.G. Chapman, 1 ♂ (OBS); Geauga Co., locality unknown, 18 July 1972, A. V. Provonsha & E. Stebbins Gulch, 18 June 1953, A.R. Gaufin, 1 ♂ (BYU); Levine, 1 ♂ (PU); Harrison Co., Buck Creek, 10 mi SE Greene Co., Little Miami River, Clifton Falls, 19 June Corydon, 9 June 2000, S. A. Grubbs, 3 ♂ (WKU); 1952, A.R. Gaufin, 3 ♂ (BYU); Highland Co., Big Jackson Co., Guthrie Creek, Hemlock Bluffs Nature Branch, 4 km NW New Petersburg, 26 June 1999, S. Preserve, 18 km E Bedford, 10 June 2000, S. A. Cordas & J. Thompson, 2 ♂ (OBS); Bull Creek, C.R. Grubbs, 5 ♂, 4 ♀ (WKU); same but 18 July 2000, S. A. 15, 1.5 km N Bridges, 26 June 1999, S. Chordas & J. Grubbs, 4 ♂, 9 ♀ (WKU); Salt Creek, nr. Maumee, 22 Thompson, 25 ♂, 11 ♀ (OBS); Jefferson Co., Brush June 2008, R.E. DeWalt, 2 ♂ (INHS); Jefferson Co., Creek, C.R. 72, 14 km SW East Liverpool, 26 June Clifty Falls State Park, 5 July 1937, collector 2000, E.G. Chapman, 2 ♂, 5 ♀ (OBS); Logan Co., Lee unknown, 1 ♂, 4 ♀ (PU); Orange Co., Patoka River, Creek, T.R. 69, 25 July 1994, V. Fazio, 1 ♂, 3 ♀ (OBS); Hoosier National Forest, 11 km SE Paoli, 15–19 June Miami Co., Spring Creek, 7 June 1953, A.R. Gaufin, 5 2006, S.A. Grubbs, 3 ♂, 2 ♀, 9 nymphs (WKU); Owen ♂ (BYU); Portage Co., Silver Creek, 3 km NW Co., Raccoon Creek, Green Bluffs Nature Preserve, 3 Garrettsville, 16 June 1953, A.R. Gaufin, 1 ♂ (BYU). km NW Freeman, 7–11 June 2006, S.A. Grubbs, 3 ♂, 6 Shelby Co., Tawawa Creek, 9 July 1994, Kovarik, 2 ♂, ♀, 20 nymphs (WKU); McCormick’s Creek, Spencer, 4 ♀ (OBS); Plum Creek, at confluence with Miami 7 June 1944, W. Stehr, 2 ♂, 1 ♀ (CNC). KENTUCKY: River, Sidney, 28 July 1970, M.A. Tkac, 1 ♂ (OBS); Green Co., Upper Brush Creek, 14 km SE Buffalo, 21 Tuscarawas Co., small woodland stream, 6 July 1997, July 2003, S.A. Grubbs, 2 ♂, 1 ♀ (WKU); same but 1 E Chapman, 1 ♂, 18 ♀ (OBS); . PENNSYLVANIA: July 2005, S.A. Grubbs, 1 ♂ (WKU); same but 17 July Crawford Co., Linesville Creek, 3 km N Linesville, 1- 2006, S.A. Grubbs, 1 ♂, 2 ♀ (WKU); Taylor Co., 5 July 1998, S.A. Grubbs, 3 ♂, 1 ♀ (WKU); same but Middle Pitman Creek, 5 km NW Campbellsville, 5 28 June 2007, S.A. Grubbs, 3 ♂, 3 ♀ (WKU); Erie Co., June 2006, S.A. Grubbs, 2 ♂ (WKU); Warren Co., Big West Branch Conneaut Creek, 11 km NW Reedy Creek, 19 km N Richardsville, 19 June 2001, Springboro, 27 June 2007, S.A. Grubbs, 1♂ (WKU). S.A. Grubbs, 1 ♂ (WKU); Doty Creek, 11 km SW Bon TENNESSEE: Grundy Co., Savage Creek, Rte. 399, 8 Ayr, 24 June 2001, S.A. Grubbs, 1 ♂ (WKU); same but km NW Dunlap, 29 May 2003, S.A. Grubbs, 1 ♂, 1 ♀, 6 July 2003, S.A. Grubbs, 2 ♂, 3 ♀ (WKU). 5 nymphs (WKU); same but 30 May 2006, S.A. MARYLAND: Allegany Co., Fifteenmile Creek, Grubbs, 1 ♀, 4 nymphs (WKU). VIRGINIA: Green Ridge State Forest, 8 km NW Little Orleans, 4 Montgomery Co., tributary to Mill Creek, Rte. 8, July 1996, S.A. Grubbs, 13 ♂, 14 ♀ (WKU); Flintstone nymphs collected 25 June 1996, adults emerged 30 Creek, Flintstone, 29 July 1996, S.A. Grubbs, 1 ♂, 3 ♀ June 1996 (reared), S. Hiner, 2 ♂, 3 ♀ (CSU); Wythe (WKU); same but 4 July 1996, S.A. Grubbs, 18 ♂, 21 ♀ Co., Reed Creek, U.S. 11 bridge, 23 July 1996, B.C. (WKU); Sideling Hill Creek, nr. Bellegrove, at light, 4 Kondratieff & R.F. Kirchner, 5 ♀, 1 nymph (Paratypes July 1996, S.A. Grubbs, 1 ♂, 1 ♀ (WKU); same but 6 – P. teaysia; CSU); Stony Fork, Reed Creek, U.S. 52, August 1996, S.A. Grubbs, 1 ♂ (WKU); same but 14 11.5 km NW Wytheville, Jefferson National Forest July 1998, S.A. Grubbs, 1 ♂ (WKU); Town Creek, (JNF), 24 July 2008, S.A. Grubbs, 3 ♀ (WKU); Reed Green Ridge State Forest, 5 km NE Oldtown, 4 July Creek, F.R. 221, 21 km WNW Wytheville, JNF, 24 July 1996, S.A. Grubbs, 3 ♂ (WKU). OHIO: Ashland Co., 2008, S.A. Grubbs, 1 ♀ (WKU); Reed Creek, C.R. 658, Jerome Fork, Mohican River, C.R. 1302, 26 June 2000, 5 km SW Wytheville, 24 July 2008, S.A. Grubbs, 2 ♀ E. Chapman, 2 ♂ (OBS); Lang Creek, US-250/OH-60, (WKU). 5 km NNW Ashland, 6 July 1995, collector unknown, Remarks. Perlesta teaysia was described from 7 ♂, 8 ♀ (OBS); Athens Co., Athens, 23 July 1997, G. southwestern Virginia (Kirchner & Kondratieff,

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Grubbs, S.A. & R.E. DeWalt 2008. Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee Plecoptera: Perlidae). Illiesia, 4(14):143-149. Available online: http://www2.pms-lj.si/illiesia/Illiesia04-14.pdf

1997). DeWalt (2002) described P. napacola from characteristics and chorionic sculpturing pattern eastern Illinois, yet first reported this species as P. nr. were identical for all eggs examined (Figs. 1–15), teaysia in DeWalt et al. (2001). DeWalt (2002) revealing that the egg depicted in Kirchner & indicated that P. napacola was distinct from P. teaysia Kondratieff (1997; Fig. 10) does not represent the true only in the egg stage, noting that males and females P. teaysia and that P. napacola must be a junior of the two species were morphologically inseparable. synonym. The combination of (a) a long tubular Perlesta teaysia was later reported as new to Maryland aedeagus lacking a caecum, (b) dense, separated (Grubbs & Stark, 2001), Pennsylvania (Grubbs & patches of sensilla basiconica on the 10th tergum, and Stark, 2001) and West Virginia (Tarter & Nelson, (c) paraprocts lacking a spine is now considered 2006) based solely on male morphology. Stark (2004), distinctive for the male of P. teaysia. in his key to the Perlidae of the eastern Nearctic The egg presented in the original description region, separated the males of the two species only belongs to an undetermined species. In addition, it by geographic differences (i.e., Illinois vs. Virginia). should be noted that the egg shown in Stark (2004; Grubbs (2004) and Tarter et al. (2006) have Fig. 7.407) for P. teaysia also is of undetermined subsequently reported P. napacola from Indiana and identity. All previously documented records of P. Kentucky, respectively, based on SEM work with napacola from Illinois (DeWalt et al., 2001; DeWalt, eggs. 2002), Indiana (Grubbs, 2004), and Kentucky (Tarter Eggs from multiple series of P. teaysia collections et al., 2006) are now referable to P. teaysia, while the from western Maryland were examined recently with collection from Tennessee represents a new state SEM, surprisingly revealing an identical pattern of record. chorionic sculpturing to P. napacola eggs, yet also very distinct from P. teaysia. This discrepancy raised Perlesta golconda DeWalt & Stark a question regarding the true identity of P. teaysia (Figs. 16–17) because the Maryland location and the Virginia type locality both lay within the Ridge and Valley Perlesta golconda DeWalt et al., 1998:315. Holotype ♂ Physiographic Province. (Illinois Natural History Survey). Ohio River, Pope Co., Eggs from paratype females of P. napacola were Illinois, U.S.A. examined with SEM and compared to eggs from Perlesta golconda Stark, 2004:93 Indiana, Kentucky, and Maryland. Eggs were not present in the type series of P. teaysia, yet gravid Material examined. U.S.A., ILLINOIS: Carroll Co., females were collected in July 2008 from each of three Savanna, at light, 20 July 1927, T.H. Frison & R.D. sites near the type locality. The egg shape, collar Glasgow, 1 ♂, 3 ♀ (INHS).

Figs. 16–17. Perlesta golconda egg SEM micrographs, U.S.A., IL, Carroll Co., Savanna, 20 July 1927. 16. Entire egg, 200X, 17. Anterior pole, 1500X.

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Grubbs, S.A. & R.E. DeWalt 2008. Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee Plecoptera: Perlidae). Illiesia, 4(14):143-149. Available online: http://www2.pms-lj.si/illiesia/Illiesia04-14.pdf

Remarks. This species was described from the Ohio Wythe Co., Stony Fork, Reed Creek, U.S. 52, 11.5 km River in southeastern Illinois (DeWalt et al., 1998). NW Wytheville, Jefferson National Forest (JNF), 24 The known distribution of this species has since been July 2008, S.A. Grubbs, 1 ♂ (WKU); Reed Creek, F.R. expanded to Iowa (Stark et al., 2008), Missouri (Stark, 221, 21 km WNW Wytheville, JNF, 24 July 2008, S.A. 2004) and Nebraska (DeWalt et al., 2001). Females Grubbs, 2 ♂ (WKU); Reed Creek, C.R. 658, 5 km SW from the original type series possessed eggs that Wytheville, 24 July 2008, S.A. Grubbs, 1 ♂ (WKU). were too poorly sclerotized to permit SEM work and Remarks. This species was described from the because only line illustrations were presented, fine unglaciated southern portions of Illinois and Indiana details of the collar and chorion could be interpreted (Grubbs, 2005). The male is distinctive among Perlesta solely from the written description (DeWalt et al., and readily identified by the combination of (a) a 1998; Fig. 8). Material stored at INHS that has been long, tubular aedeagus lacking a caecum, (b) dense subsequently identified as P. golconda included separated patches of sensilla basiconica on the 10th gravid females, allowing for SEM and the first high- tergum, and (c) long slender paraprocts bearing a resolution images of the eggs of this species. laterally-directed tooth. As part of a larger study Although the eggs depicted here were too fragile to addressing elevational patterns of stoneflies of the clean sufficiently, they adhere easily to the original montane Talladega National Forest region of east- description in DeWalt et al. (1998). All eggs examined central Alabama, Perlesta specimens collected in were near-symmetrically oval, the chorion was summer 2006 included males that were tentatively smooth to slightly granular, and the collar was raised identified as P. shawnee. Because mature eggs were only slightly above the anterior surface of the egg not present in females, however, a definitive species (Figs. 16–17). designation was not applied. Gravid females were collected in July 2008 and Perlesta shawnee Grubbs eggs were examined via SEM. Although eggs from (Figs. 18–23) the Alabama series (Fig. 19) were noticeably less pitted and smoother than from paratype females Perlesta shawnee Grubbs, 2005:64. Holotype ♂ (Illinois from Indiana and Illinois (Figs. 21, 23), both the Natural History Survey). Cooper Creek, Alexander Co., overall egg shape (i.e., oval; Figs. 18, 20, 22) and Illinois, U.S.A. Perlesta shawnee Stark, 2004:95 collar (i.e., low, gently-rounded open pit; Figs. 19, 21, 23) were too similar to warrant a new species Material examined. U.S.A., ALABAMA: Clay Co., description. tributary to Cheaha Creek, Turnipseed Campground, In addition, several series of this species were also Talladega National Forest (TNF), 11 July 2008, S.A. obtained recently from southwestern Virginia and Grubbs, 1 ♀ (WKU); Swept Creek, F.R. 615, 20 km S western North Carolina. The occurrence of P. shawnee Talladega, TNF, 20 June 2006, A.L. Sheldon, 1 ♂ in Alabama, North Carolina and Virginia represent (WKU); tributary to Swept Creek, F.R. 615, TNF, 25 new state records and a significant southeastern May 2006, A.L. Sheldon, 1 ♀ (WKU); Cleburne Co., range extension from Illinois and Indiana. tributary to South Fork Terrapin Creek, F.R. 500, 8 km SE Piedmont, TNF, 11 July 2008, S.A. Grubbs, 2 ♀ ACKNOWLEDGEMENTS (WKU); Talladega Co., Smelley Creek, C.R. 234, 15 Paul Tinerella (Illinois Natural History Survey) km S Talladega, TNF, 26 May 2006, A.L. Sheldon, 1 ♂ and Boris Kondratieff (Colorado State University) (WKU). ILLINOIS: Pope Co., Quarrel Creek. 8 km NE made the paratypes of Perlesta napacola and P. teaysia, Dixon Springs, 13 July 2003, S.A. Grubbs 2 ♂, 3 ♀ respectively, available for study. Boris Kondratieff (Paratypes; WKU); Lusk Creek, 13 km NW Golconda, kindly provided the North Carolina record of P. 20 May 2006, S.A. Grubbs 1 ♂ (WKU). INDIANA: shawnee. Brian Armitage (Ohio Biological Survey), Perry Co., East Deer Creek, 13 km E Tell City, Shawn Clark and Richard Baumann (Brigham Young Hoosier National Forest, 12 June 2003, S.A. Grubbs 1 University), Arwin Provonsha (Purdue University) ♂, 2 ♀ (Paratypes; WKU). NORTH CAROLINA: and Ian Smith (Canadian National Collection) either Cleveland Co., First Broad River, Mt. Zion Church provided loans or accommodated visits to examine Rd., S of New Hope, 13 July 2007, B.C. Kondratieff, material onsite. John Andersland (Western Kentucky R.E. Zuellig & D.R. Lenat 1 ♂, 6 ♀ (CSU). VIRGINIA: University) assisted with SEM work.

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Grubbs, S.A. & R.E. DeWalt 2008. Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee Plecoptera: Perlidae). Illiesia, 4(14):143-149. Available online: http://www2.pms-lj.si/illiesia/Illiesia04-14.pdf

Figs. 18–23. Perlesta shawnee egg SEM micrographs. 18–19: U.S.A., AL, Cleburne Co., tributary to South Fork Terrapin Creek, 11 July 2008, 20-21: U.S.A., IL, Pope Co., Quarrel Creek, 13 July 2003, 22-23: U.S.A., IN, Perry Co., East Deer Creek, 12 June 2003), 18. Entire egg, 350X, 20, 22. Entire egg, 200X, 19, 21, 23. Anterior pole, 2000X.

REFERENCES the Perlesta frisoni species group (Plecoptera: DeWalt, R.E. 2002. Perlesta napacola, a new species in Perlidae). Entomological News, 113:173–178.

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Grubbs, S.A. & R.E. DeWalt 2008. Taxonomic and distributional notes on Perlesta teaysia, P. golconda, and P. shawnee Plecoptera: Perlidae). Illiesia, 4(14):143-149. Available online: http://www2.pms-lj.si/illiesia/Illiesia04-14.pdf

DeWalt, R.E., B.P. Stark, & M.A. Harris. 1998. Perlesta (USA). Proceedings of the Entomological Society golconda (Plecoptera: Perlidae), a new stonefly of Washington, 108:429–442. species from Illinois. Entomological News, Tarter, D.C., D.L. Chaffee, & S.A. Grubbs. 2006. 109:315–317. Revised checklist of the stoneflies (Plecoptera) of DeWalt, R.E., D.W. Webb, & T.N. Kompare. 2001. Kentucky, U.S.A. Entomological News, 117:1–10. The Perlesta placida (Plecoptera: Perlidae) complex in Illinois: new state records, distributions, and an identification key. Proceedings of the Entomological Society of Washington, 103:207– Received 23 September 2008, Accepted 17 October 2008, Published 216. 29 October 2008 Grubbs, S.A. 2004. Studies on Indiana stoneflies (Plecoptera), with an annotated and revised state checklist. Proceedings of the Entomological Society of Washington, 106:865–876. Grubbs, S.A. 2005. Perlesta shawnee (Plecoptera: Perlidae), a new stonefly species from eastern North America. Aquatic , 27:63–69. Grubbs, S.A. & B.P. Stark. 2001. Notes on Perlesta (Plecoptera: Perlidae) from eastern North America. Aquatic Insects, 23:119–122. Kirchner, R.F. & B.C. Kondratieff. 1997. A new species of Perlesta (Plecoptera: Perlidae) from Virginia. Proceedings of the Entomological Society of Washington, 99:290–293. Kondratieff, B.C., R.E. Zuellig, R.F. Kirchner, & D.R. Lenat. 2006. Three new species of Perlesta (Plecoptera: Perlidae) from eastern North America and notes on new state records. Illiesia, 2:31–38 Kondratieff, B.C., R.E. Zuellig, R.F. Kirchner, & D.R. Lenat. 2008. Two new species of Perlesta (Plecoptera: Perlidae) from eastern North America. Proceedings of the Entomological Society of Washington, 110:668–673. Stark, B.P. 1989. Perlesta placida (Hagen), an eastern Nearctic species complex (Plecoptera: Perlidae). Entomologica Scandinavica, 20:263–286. Stark, B.P. 2004. Perlidae (The Stones). Pages: 61–148. In B.P. Stark & B.J. Armitage, editors. Stoneflies (Plecoptera) of Eastern North America. Volume II. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). The Caddis Press, Columbus, OH, U.S.A., 192 pp. Stark, B.P., R.W. Baumann & R.E. DeWalt. 2008. Valid stonefly names for North America. Updated 28 January 2008. HTML address, http://plsa.inhs.uiuc.edu/plecoptera/validnames.a spx. Accessed July 2008. Tarter, D.C. & C.H. Nelson. 2006. A revised checklist of the stoneflies (Plecoptera) of West Virginia

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